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Kunming Institute of Botany, Chinese Academy of Sciences, Heilongtan, Kunming 650204, P. R. China
M. Weiß
F. Oberwinkler
Botanisches Institut, Spezielle Botanik und Mykologie, Auf der Morgenstelle 1, D-72076 Tübingen, Germany
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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Four new species of Amanita, Amanita-ceae (Agaricales) are described from the eastern Himalaya and adjacent regions of southwestern China. Amanita altipes and A. parvipantherina are members of section Amanita, while A. orientifulva and A. liquii are representatives of section Vaginatae. They are compared with similar species and illustrated with line drawings.
Key words: Amanitaceae, Agaricales, Basidiomycetes, China, new taxa, systematics
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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; color codes of the form ‘‘3A4’’ that indicates the plate, row, and color block are from Kornerup and Wanscher (1981). Herbarium codes used follow Holmgren et al (1990) with one exception: HKAS = Herbarium of Cryptogams, Kunming Institute of Botany, Chinese Academy of Sciences, which is not listed in the Index Herbariorum or relative publications. |
TAXONOMY |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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Etymology: altipes, referring to the long stipe.
Basidiocarps (FIG. 1
) small to medium-size. Pileus 4–9 cm diam, convex to planoconvex, broadly umbonate, yellowish to yellow (Apricot Yellow, Colonial Buff, Mustard Yellow, Straw Yellow, Warm Buff, 3A4–8), often with brownish tinge (Cinnamon-Buff, Orange-Cinnamon, 5C6–8, 5D6–8) over disk, viscid when wet; margin tuberculatestriate (0.13–0.41R), pale yellow (Pale Orange-Yellow, Light Orange-Yellow, 3A2–5), nonappendiculate; volval remnants as felty, floccose patches, 2–5 mm wide, up to 1 mm thick, yellowish to yellow to dirty yellow (Colonial Buff, Maize Yellow, Buff-Yellow, 3A2–5), randomly arranged, often washed away by rain or retained in the humus; trama white, unchanging. Lamellae free, crowded, white to cream-colored to yellowish (Cartridge Buff, Cream Color, Maize Yellow, 2A1–4), with yellowish to yellow edges; lamellulae truncate, plentiful, evenly distributed. Stipe 9–16 x 0.5–1.8 cm, subcylindric or tapering upward, with apex slightly expanded, yellowish (Straw Yellow, Light Orange-Yellow, 2A2–4, 3A2–4), but becoming whitish toward stipe base, covered with yellow to yellowish (Apricot Yellow, Mustard Yellow, Straw Yellow, 3A4–8) squamules above annulus, with yellowish to whitish squamules or fibrils under annulus; context white, hollow in center; basal bulb subglobose to ovate, 0.8–3.2 cm diam, white to pallid; volval remnants as yellow to yellowish (Apricot Yellow, Mustard Yellow, Straw Yellow, 3A6–8) floccose patches or warts near apex of bulb, occasionally forming a short limb. Annulus persistent, pendant from attachment 1–3 cm below apex of stipe, upper surface cream-colored to yellowish (Cream Color, paler than Maize Yellow, 2A2–4), lower surface yellowish, with a yellow edge. Odor indistinct.
Lamellar trama bilateral. Mediostratum 50–70 µm wide, composed of abundant fusiform to subellipsoid inflated cells (70–90 x 15–30 µm), mixed with abundant, 2–6(–8) µm wide filamentous hyphae, vascular hyphae rare. Lateral stratum composed of abundant clavate to subellipsoid inflated cells (50–90 x 10–25 µm), diverging at an angle of ca 30–45° to the mediostratum; filamentous hyphae abundant, 3–7 µm wide; septa without clamps. Subhymenium (FIG. 2) 20–40 µm thick, with 2–3 layers of ovoid to subellipsoid to irregularly formed cells, 9–25 x 8–20 µm. Basidia (FIG. 2) (35–)40–60(–75) x 10–16 µm, clavate, 4-spored, rarely 2-spored; sterigmata 3–6(–8) µm long; basal septa without clamps. Basidiospores (FIG. 3) (363/16/7) (7.5–)8.0–10.0(–11.5) x (7.0–) 7.5–9.5(–10.0) µm (Q = 1.0–1.14[–1.20], Q = 1.07 ± 0.04), globose to subglobose, rarely broadly ellipsoid, inamyloid, colorless, hyaline, thin-walled, smooth; apiculus small. Lamellar edge appearing as sterile, somewhat gelatinized strip 100–150 µm wide in side view, composed of numerous clavate to broadly clavate (30–40 x 10–20 µm) or ovoid to subglobose (18–25 x 14–20 µm) to ellipsoid (18–25 x 8–15 µm) inflated cells, single and terminal or in chains of 2–3, thin-walled, colorless, hyaline or with yellowish contents; filamentous hyphae abundant, 2–5 µm wide, thin-walled, hyaline or with yellowish vacuolar pigments, ± running parallel to the lamellar edge. Pileipellis 100–150 µm thick; upper layer (30–100 µm thick) strongly gelatinized, composed of subradially arranged, 2–3(–5) µm wide, thin-walled, colorless to subcolorless, filamentous hyphae, often with fine granular incrustations; lower layer (50–70 µm thick) composed of radially and compactly arranged, 3–5 µm wide hyphae, subcolorless or with yellowish vacuolar pigments; vascular hyphae rare, 3–7 µm wide. Volval remnants on pileus (FIG. 4) composed of ± radially to somewhat irregularly arranged elements; filamentous hyphae fairly abundant to abundant, becoming more abundant toward the pileal surface, 2–7 µm wide, thin-walled, colorless, hyaline, branching, interwoven, sometimes anastomosing; inflated cells abundant to very abundant, ovoid to subglobose (35–60 x 30–55 µm) to broadly ellipsoid (40–65 x 25–35 µm) to long ellipsoid (45–90 x 15–35 µm), sometimes broadly clavate (60–70 x 28–35 µm), single and terminal or in chains of 2–3 and then terminal, thin- to slightly thick-walled (up to 1 µm thick), colorless to subcolorless; vascular hyphae rare. Outer surface of volval remnants a layer of 10–30 µm thick composed of gelatinized and collapsed yellowish to brownish filamentous hyphae, mixed with collapsed or uncollapsed inflated cells similar to those in the interior. Volval remnants on the stipe base composed of ± irregularly arranged elements; filamentous hyphae abundant to very abundant to locally nearly dominant, 2–7(–15) µm wide, thin- to slightly thick-walled (ca 0.5 µm thick), colorless, hyaline, sometimes with yellowish vacuolar pigments; inflated cells abundant to locally very abundant, subglobose to ovoid (20–60 x 15–50 µm) to ellipsoid (30–90 x 12–35 µm), single and terminal or in chains of 2–3 and then terminal, colorless, hyaline, occasionally with yellowish vacuolar pigments, thin-to slightly thick-walled (up to 1 µm thick); vascular hyphae rare. Stipe trama composed primarily of longitudinally arranged, long clavate, terminal cells, 260–450 x 25–40 µm; filamentous hyphae scattered (in interior) to fairly abundant (on stipe surface), 2–7(–12) µm wide; vascular hyphae rare, 3–10 µm wide. Annulus composed of loosely and subradially arranged, 2–5(–8) µm wide, thin-walled, anastomosing, filamentous hyphae; inflated cells scattered to locally fairly abundant, clavate to broadly clavate to ellipsoid (35–70 x 10–25 µm), thin-walled, hyaline, usually single and terminal; vascular hyphae rare. Upper surface of annulus gelatinized, elements often with yellow to yellowish vacuolar pigments and incrustations.
Habit, habitat and distribution. – Solitary to scattered on soil under Abies, Picea, Quercus, Betula and/or Salix; fruiting from August to September in southwestern China at 2700–4000 m elevation.
Specimens examined..
CHINA. SICHUAN PROVINCE: Hongyuan County, Kangle, altitude 3400 m, 11 Aug 1998, M. S. Yuan 3263 (HKAS 33847). Luding County, Moxi, altitude 3200 m, 23 Aug 1997, D. S. Hibbett 97–183 (FH). XIZANG AUTONOMOUS PREFECTURE (TIBET): Bomi County, Guxiang, altitude 2700 m, 12 Sep 1982, X. L. Mao 419 (HMAS 52601, as A. citrina in Mao 1990). YUNNAN PROVINCE: Lijiang County, Laojunshan, altitude 3800 m, 14 Aug 2000, Zhu L. Yang 2915 (HKAS 36609, HOLO-TY PE). Zhongdian County, Daxueshan, altitude 4000 m, 23 Aug 2000, Zhu L. Yang 2992 (HKAS 36596); same locality, 24 Aug 2000, Zhu L. Yang 3007 (HKAS 36606). Zhongdian County, Tianchi, altitude 3900 m, 20 Aug 2000, Zhu L. Yang 2951 (HKAS 36590).
Commentary. –
Amanita altipes, belonging in Amanita subgenus Amanita section Amanita (Yang 1997), resembles A. orientigemmata Zhu L. Yang et Yoshim. Doi and A. pantherina var. lutea W. F. Chiu, both originally described from East Asia. However, A. altipes differs from A. orientigemmata by its more slender basidiocarps with a superior annulus, wider basidiospores and radially to somewhat irregularly arranged elements with rare vascular hyphae in the volval remnants on the pileus, and absence of clamps (Yang and Doi 1999). The basidiospores of A. orientigemmata are (7.5–)8.0–10.0(–11.0) x (6.0–)6.5–7.5(–8.0) µm (Q = [1.07]1.15–1.46[–1.59], Q = 1.30 ± 0.01) (Yang and Doi 1999). Amanita pantherina var. lutea is distinguished from A. altipes by its whitish to grayish warts on the pileus, paler-colored gills, and proportionately narrower basidiospores with higher Q ratio. Chiu (1948) described the basidiospores of A. pantherina var. lutea as (8)9–10 x 6–7 µm, and Yang (1997) reported the basidiospores are (8.0)8.5–11.5(12.5) x (6.0)7.0–8.5(9.5) µm (Q = [1.06]1.13–1.47[–1.53], and Q = 1.29 ± 0.11). Furthermore, the warts on the pileus of A. pantherina var. lutea are composed of more or less vertically arranged filamentous hyphae and inflated cells (Yang 1997).
Species in the section Amanita having yellow to yellowish pilei and exhibiting some morphological similarities to A. altipes are A. elata (Massee) Corner et Bas, A. gemmata (Fr.) Bertillon, A. russuloides (Peck) Sacc., A. xylinivolva Tulloss, Ovrebo et Halling. However, A. elata, originally described from Singapore, differs from A. altipes by its more or less umber or fuliginous pileal disk, exannulate stipe, an unpleasant odor and smaller basidiospores (Corner and Bas 1962). The basidiospores of A. elata are 7.0–8.5 x (6.0)6.8–7.7 µm (Q = 1.0–1.1[–1.2]) (Corner and Bas 1962). Amanita gemmata, originally described from Europe, differs from A. altipes by its relatively shorter stipe, white volval remnants with fewer inflated cells and basidiospores with higher Q ratio. Breitenbach and Kränzlin (1995) reported the basidiospores of A. gemmata are 8.9–10.8 x 6.8–8.7 µm (Q = 1.1–1.4). Yang and Doi (1999) reported the spores of A. gemmata are (9.0–)9.5–12.0(–13.0) x (6.0–) 6.5–8.5(–9.0) µm (Q = [1.17]1.25–1.62[–1.77], Q = 1.42 ± 0.13). Amanita russuloides (Peck) Sacc., originally described from North America, differs from A. altipes by its white, warty volval remnants on the pileus, white stipe and narrower basidiospores ( Jenkins 1977, Tulloss et al 1995). The basidiospores of A. russuloides are 8.7–10.2 x 6.3–7.0 µm according to Jenkins (1977). Tulloss et al (1995) reported the basidiospores of A. russuloides are (8.7)9.8–10.5(10.8) x 6.6–7.7 µm. Amanita xylinivolva, originally described from Colombia, has fruit bodies without a pileal umbo, white to off-white to dingy gray volval remnants, whitish buff lamellae and marked to abrupt bulb (Tulloss et al 1992).
In China A. altipes probably was regarded as A. pantherina (DC. : Fr.) Krombh. originally described from Europe. But the latter is distinguished from the former by its brownish to brown pileus with white verrucous to subconical volval remnants, a proportionately shorter stipe and broadly ellipsoid to ellipsoid basidiospores. Breitenbach and Kränzlin (1995) reported the basidiospores of A. pantherina are 8.9–11.5 x 7.0–8.4 µm (Q = 1.2–1.5). In addition, in A. pantherina the volval remnants on the pileus are composed of vertically arranged elements (unpublished data of Yang studying material from the Netherlands, Bas 7474 [L]). HMAS 52601 was regarded incorrectly as A. citrina (Schaeff.) Pers. by Mao (1990), but the European A. citrina has, among other features, a nonstriate pileus, a subtruncate bulb on the stipe base and amyloid basidiospores (Breitenbach and Kränzlin 1995).
Amanita parvipantherina Zhu L. Yang, M. Weiss et Oberw., sp. nov. FIGS. 5–8
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Etymology: parvipantherina, a smaller mushroom similar to A. pantherina.
Basidiocarps (FIG. 5) small to medium-size. Pileus 3.5–6 cm diam, convex to planoconvex, often slightly depressed at center, grayish to ochraceous (Light Drab, Buffy Brown, 5B2–3, 5C4–6), becoming brown to brownish (Snuff Brown, Cinnamon-Brown, 5D4–6, 5E6–7) over disk, viscid when moist; margin tuberculate-striate (0.27–0.4R), nonappendiculate; volval remnants as conical, subconical to granular, small warts, up to 2 mm high, dirty white, grayish to yellowish (Cream Color, Pale Smoke Gray, Cream Buff, 2A1–3, 5B2), randomly arranged, frequently densely placed over disk; trama white, unchanging. Lamellae free, white (1A1), crowded, with finely fimbriate and floccose edges; lamellulae truncate, plentiful, in 2–3 ranks. Stipe 4–9 x 0.5–1 cm, subcylindric or slightly tapering upward, with apex slightly expanded, white (1A1), fibrillose to subglabrous; context white, loosely stuffed to hollow in center; basal bulb 1–2 cm diam, subglobose to napiform, white to pallid, with upper part covered with floccose to granular, dirty white to yellowish or grayish (Cream Color, Pale Smoke Gray, Cream Buff, 2A1–3, 5B2) volval remnants. Annulus present, superior, pendant from attachment 1.5–2.5 cm below apex of stipe, white (1A1), thin, membranous. Odor indistinct.
Lamellar trama bilateral. Mediostratum 30–40 µm wide, composed of long ellipsoid to subfusiform, inflated cells, 90–130 x 10–30 µm; filamentous hyphae fairly abundant, 2–7 µm wide; vascular hyphae rare. Lateral stratum composed of long ellipsoid to fusiform inflated cells, 55–100 x 20–30 µm, diverging at an angle of 30–45° to the mediostratum; filamentous hyphae fairly abundant, 3–7 µm wide; septa without clamps. Subhymenium (FIG. 6) 30–40 µm thick, with 2–3(–4) layers of subglobose, ovoid to barrel-shaped cells, 10–20 x 8–17 µm. Basidia (FIG. 6) 38–55 x 10–13 µm, clavate, 4-spored; sterigmata 3–5 µm long; basal septa without clamps. Basidiospores (FIG. 7) (360/14/7) (8.0–)8.5–11.5(–13.5) x (6.0–)6.5–8.5 (–9.0) µm (Q = [1.13–]1.22–1.54[–1.62], Q = 1.38 ± 0.10), broadly ellipsoid to ellipsoid, rarely subglobose or elongate, inamyloid, colorless, hyaline, thin-walled, smooth; apiculus small. Lamellar edge appearing as sterile strip up to 100 µm wide in side view, composed of clavate to broadly clavate to sphaeropedunculate cells (25–40 x 10–22 µm), single and terminal or in terminal chains of 2–3, thin-walled, colorless, hyaline; filamentous hyphae very abundant, 2–5 µm wide, gelatinized, irregularly arranged or ± running parallel to the lamellar edge. Pileipellis 100–150 µm thick; upper layer (60–80 µm thick) strongly gelatinized, composed of subradially arranged to somewhat interwoven 3–5 µm wide, colorless to sub-colorless, moderately gelatinized, filamentous hyphae; lower layer (40–70 µm thick) composed of radially and compactly arranged, 3–6 µm wide filamentous hyphae, sometimes with intercalary segments inflated to 8–10(–12) µm wide, often with brownish vacuolar pigments; vascular hyphae rare, 3–6 µm wide. Volval remnants on pileus (FIG. 8) composed of ± vertically, at wart base subvertically, arranged elements; inflated cells abundant to nearly dominant, subglobose, ovoid to short ellipsoid (20–65 x 10–40 µm) or ellipsoid to subfusiform (35–60 x 15–30 µm), often in chains of 2–4, thin-walled to slightly thick-walled (ca 0.5 µm thick), colorless, hyaline to subhyaline, sometimes with fine incrustations; filamentous hyphae fairly abundant, 2–8 µm wide, thin-walled, colorless, hyaline to subhyaline; vascular hyphae rare or locally conspicuous, 2–7 µm wide. Volval remnants on stipe base and upper bulb with structure similar to those on pileus, but elements irregularly arranged and filamentous hyphae more abundant. Stipe trama composed primarily of longitudinally arranged, long clavate, terminal cells, 200–410 x 25–45 µm; filamentous hyphae 2–7(–10) µm wide, scattered to fairly abundant in interior, abundant on stipe surface; vascular hyphae rare. Annulus composed primarily of filamentous, 2–7 µm wide hyphae, somewhat loosely and subradially arranged, frequently branching, anastomosing, interwoven, thin-walled, colorless, hyaline; inflated cells fairly abundant to locally abundant, clavate to long ellipsoid (35–130 x 10–30 µm), occasionally subglobose to ovoid (20–45 x 15–30 µm), usually single and terminal, rarely 2–3 in chains; vascular hyphae rare.
Habit, habitat and distribution. – Solitary to scattered on soil in mixed forests with Pinus yunnanensis; fruiting from July to August in Yunnan, southwestern China at 1200–2500 m elevation.
Specimens examined.. CHINA. YUNNAN PROVINCE: Binchuan County, Jizushan, 9 Aug 1985, G. P. Xiao 529 (HKAS 17032). Heqing County, Huangping, altitude 2500 m, 24 Jul 1998, Zhu L. Yang 2490 (HKAS 32492, HOLOTY PE). Menghai County, Mangao Nature Reserve, altitude 1200 m, 14 Aug 1991, Zhu L. Yang 1519 (HKAS 24180). Simao County, Zhala Yakou, altitude 1730 m, 1 Aug 1991, Zhu L. Yang 1338 (HKAS 24202). Tengchong County, Mingguang, 2 Aug 1980, M. Zang 6301 (HKAS 6301). Wuding County, Sishan, 27 Jul 1988, L. S. Wang 9 and 10 (HKAS 20886 and 20887 respectively).
Commentary. –
Amanita parvipantherina is a member of Amanita subgenus Amanita section Amanita (Yang 1997). A few collections cited above were regarded incorrectly as A. pantherina (DC. : Fr.) Krombh., but the latter has more robust basidiocarps with shorter striations on the pileal margin, a short limbate collar on the stipe base and more abundant filamentous hyphae in the white volval remnants on the pileus (Breitenbach and Kränzlin 1995; Yang 1997, unpublished data of Yang studying material from the Netherlands (THE NETHERLANDS. ZÜID-HOLLAND: Wassenaar, 2 Sep 1979, C. Bas 7474 [L]). Amanita parvipantherina is similar to A. sychnopyramis f. subannulata Hongo (Hongo 1971) but differs from the latter by its somewhat smaller basidiocarps with a superior annulus and larger, broadly ellipsoid to ellipsoid basidiospores. The basidiospores of A. sychnopyramis f. subannulata are mostly globose to subglobose, (6.0–)6.5–8.5(–9.5) x 6.0–8.0(–9.0) µm (Q = [1.0]1.03–1.14[–1.19], Q = 1.07 ± 0.04) (Yang 1997). In addition, the warts on the pileus of A. parvipantherina are composed of more abundant filamentous hyphae than those of A. sychnopyramis f. subannulata. Amanita parvipantherina also resembles A. sychnopyramis Corner et Bas f. sychnopyramis but differs from the latter by the presence of an annulus, longer basidia, and larger, broadly ellipsoid to ellipsoid basidiospores. The basidia of A. sychnopyramis f. sychnopyramis are 26–32 x 10–11 µm, and the basidiospores of this taxon are globose to subglobose, 6.3–8.1 µm, Q = 1.0–1.1 (Corner and Bas 1962).
Amanita liquii Zhu. L. Yang, M. Weiss et Oberw., sp. nov. FIGS. 9–12
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Etymology: A. liquii is named for Li Qui (Li Kui), a historic Chinese hero, said to have a dark face.
Basidiocarps (FIG. 9) large. Pileus 10–14 cm diam, at first nearly hemispherical, then convex to planoconvex, center often slightly umbonate, sepia to blackish (Warm Sepia, Clove Brown, Fuscous to Fuscous-Black, 6F3–7), becoming blackish brown to dark brown (Brussels Brown, Raw Umber, Medal Bronze, Cinnamon-Brown, Natal Brown, 6D3–5, 6E3–7) toward margin, lacking any yellow tint at all stages of development; margin tuberculatestriate (0.15–0.25R), nonappendiculate; volval remnants dark gray to sepia, sometimes gray (Warm Sepia, Medal Bronze, 6D2–3, 6E2–4, 6F3–5), felty to irregularly formed to verrucous, 2–10 mm diam, 1–3 mm thick (high); trama white, but grayish to gray just beneath pileipellis, unchanging. Lamellae free, crowded, white to cream-colored (Cream Color, 1A1–2) when young, grayish (much paler than Pale Smoke Gray, paler than 3B2) when mature, becoming brownish gray, dark gray to fuscous (paler than Fuscous, 5C3–5, 5D3–5, 5E3–5) when dried; lamellar edges blackish to dark brown (Medal Bronze, Natal Brown, 6E3–5, 6F5–7); lamellulae truncate to subtruncate, plentiful, evenly distributed. Stipe 13–17 x 1.5–3 cm, subcylindric or slightly tapering upward, with apex slightly expanded, whitish to brownish, densely covered with dark gray to blackish (Warm Sepia, Medal Bronze, Raw Umber, 6E3–6, 6F4–7) squamules often in belts; context white but at very base often grayish, sometimes brownish to light rusty brown, hollow in center; basal bulb lacking; volval remnants verrucous to sub-conical to granular, gray to dark gray to brownish (Smoky Gray, Drab, Mouse Gray, 5C1–4) in apical parts, becoming paler toward base of remnants, arranged in incomplete belts at stipe base. Annulus lacking. Odor indistinct.
Lamellar trama bilateral. Mediostratum 30–50 µm wide, composed of fairly abundant, long ellipsoid to subfusiform inflated cells (30–90 x 10–25 µm) and abundant interwoven, often anastomosing, 2–7 µm wide filamentous hyphae; vascular hyphae rare. Lateral stratum composed of fairly abundant, long ellipsoid to subfusiform (55–70 x 20–28 µm), sometimes to ellipsoid (55–70 x 20–28 µm) inflated cells, diverging at an angle of ca. 30–45° to mediostratum; filamentous hyphae abundant, 2–6 µm wide, frequently branching, interwoven, sometimes anastomosing; vascular hyphae rare, 3–7 µm wide. Subhymenium (FIG. 10) 40–60(–70) µm thick, with (2)3–4 layers of subglobose to ovoid or irregularly shaped cells 12–30 x 10–25 µm, sometimes mixed with a few hardly inflated cells 5–7 µm wide. Basidia (FIG. 10) 55–95 x 16–25 µm, clavate, 4-spored, rarely 1- or 2-spored; sterigmata 6–10 µm long; basal septa without clamps. Basidiospores (FIG. 11) (220/9/7) (11.0–)11.5–15.0(–24.0) x (9.5–)11.0–14.5(–20.0) µm (Q = 1.0–1.09[–1.21], Q = 1.05 ± 0.04), globose to subglobose, rarely broadly ellipsoid, inamyloid, colorless, hyaline, thin-walled, smooth; apiculus small. Lamellar edge appearing as sterile, gray-brown to dark brown strip up to 250 µm wide in side view, predominantly composed of inflated cells, ovoid to subglobose or short ellipsoid (20–60 x 15–50 µm), often in chains of 2–3, sometimes sphaeropedunculate to pyriform (35–65 x 20–35 µm), then single and terminal, with brownish to brown to gray-brown vacuolar pigments, occasionally nearly colorless and hyaline; filamentous hyphae fairly abundant to scattered, 2–7 µm wide, with brownish vacuolar pigments or nearly colorless. Pileipellis 100–180 µm thick; upper layer (20–70 µm thick) slightly to moderately gelatinized, composed of ± radially and moderately compactly arranged, 2–5 µm wide, thin-walled, colorless to nearly colorless, slightly to moderately gelatinized filamentous hyphae; lower layer (80–120 µm thick) composed of radially and compactly arranged, 3–7 µm wide filamentous hyphae, sometimes with intercalary segments inflated to 15 µm wide, usually with dark yellowish brown, vacuolar pigments; terminal cells present, 5–12 µm wide, hardly inflated; vascular hyphae rare to locally scattered, 3–10 µm wide. Volval remnants on pileus: the verrucous remnants (FIG. 12) composed of ± vertically, at volval base irregularly, arranged elements; felty remnants composed of ± irregularly arranged elements. Inflated cells very abundant (to dominant), subglobose (25–80 x 20–70 µm) or ovoid to broadly clavate (30–50 x 25–38 µm), becoming larger toward the pileipellis, single and terminal, or in chains of 2–3(–4) and then terminal, thin- to slightly thick-walled (up to 1.0 µm thick), usually with brown to dark brown, vacuolar pigments, occasionally colorless and hyaline; filamentous hyphae fairly abundant, not becoming more abundant toward the pileipellis, 3–8 µm wide, frequently branching, often anastomosing, thin-walled, colorless, hyaline or with brownish vacuolar pigments; vascular hyphae rare, 2–8 µm wide. Volval remnants on stipe base similar to those on the pileus, but inflated cells and filamentous hyphae irregularly arranged, and the vacuolar pigments locally much paler. Stipe trama composed primarily of longitudinally arranged, clavate to long clavate, terminal cells, 70–400 x 16–50 µm; filamentous hyphae 2–7 µm wide, scattered (in interior) or fairly abundant to abundant (on stipe surface), colorless and hyaline (in interior), or with brown to dark brown vacuolar pigments (on stipe surface); vascular hyphae 3–15 µm wide, rare to locally conspicuous.
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Specimens examined.. CHINA. SICHUAN PROVINCE: Yanyuan County, Beilingshan, altitude 3820 m, 9 Aug 1983, K. K. Chen 541 (HKAS 14096, as A. ceciliae in Yang 1997). YUNNAN PROVINCE: Dali Municipality, Huadianba, altitude 2900 m, 10 Jul 1984, J. X. Xi 569 (HKAS 378, as A. ceciliae in Yang 1997). Deqin County, Baimaxueshan, altitude 3700 m, 11 Jul 1981, J. W. Chen 34 (HKAS 8575, as A. ceciliae in Yang 1997). Lijiang County, Laojunshan, altitude 3900 m, 14 Aug 2000, Zhu L. Yang 2916 (HKAS 36611, HOLOTY PE). Lijiang County, Laojunshan, altitude 4000 m, 25. Sep 2000, M. Zang 13712 (HKAS 37052). Zhong-diang County, Xiaozhongdian, altitude 3900 m, 24 Aug 1984, K. K. Chen & Y. Xuan 19 (HKAS 14242, as A. ceciliae in Yang 1997). XIZANG AUTONOMOUS PREFECTURE (TIBET): Chayu, Ridong, altitude 3500 m, 18 Sep 1982, M. Zang 1005 (HKAS 11254, as A. ceciliae in Yang 1997).
Commentary. –
Amanita liquii is characterized by its dark-colored, robust fruit body with grayish lamellae, dark-colored volval remnants on the pileus, verrucous volval remnants in belts at the nonbulbous stipe base, and globose to subglobose, inamyloid basidiospores. Several collections cited above were regarded as A. ceciliae by Yang (1997). Amanita liquii is similar to A. ceciliae (Berk. et Broome) Bas, A. beckeri Huijsman ex Huijsman, A. cinctipes Corner et Bas, and A. sororcula Tulloss, Ovrebo et Halling. Actually, several Chinese amanitas of this group have been determined as A. ceciliae, originally described from Europe (Teng 1936 and 1996 as Amanitopsis strangulata [Fr.] Roze; Mao 1990 as A. inaurata Secr. ex Gillet; Yang 1997). However, the European A. ceciliae differs from A. liquii by its yellow-brown, reddish-brown to gray-brown or olive-brown pileus covered with lighter-colored (grayish to brownish) volval remnants, white lamellae with white edge, much paler-colored squamules on the stipe and somewhat smaller basidiospores. The basidiospores of A. ceciliae are 10.4–14.1 x 9.7–14.0 µm (Breitenbach and Kränzlin 1995). Furthermore, the volval remnants at the base of the stipe of A. ceciliae often form a ring-zone above a strangulate region and a floccose, nearly cupulate structure at the very base of the stipe (Phillips 1990, Breitenbach and Kränzlin 1995). The vacuolar pigments in the cells of the sterile strip along lamellar edge and volval remnants on the pileus of A. liquii are much darker than those of European A. ceciliae (FRANCE. JURA: near Noga, 2 Oct 1992, C. Bas 9341 [L]). Amanita beckeri, originally described from Europe, differs from A. liquii by its differently colored pileus, lighter-colored volval remnants and smaller basidiospores (Huijsman 1962a, b, Tulloss 1994). The basidiospores of A. beckeri are (8.8)9.8–11.8(13.0) x (7.2)9.0–11.0(12.2) µm (Tulloss 1994). Amanita cinctipes, originally described from Southeast Asia, is distinguished from A. liquii by its slender fruit body lacking an umbo on the pileus, smaller basidia and smaller basidiospores (Corner and Bas 1962). Basidia and the basidiospores of A. cinctipes are 40–50 x 15–15 µm and 8.2–11.1 x 7.8–10.1 µm respectively (Corner and Bas 1962). Amanita sororcula, originally described from Colombia, differs from A. liquii by its smaller and more slender fruit body with longer marginal striations on the pileus, volval remnants tending to darken considerably with age and smaller basidiospores (Tulloss et al 1992, Tulloss pers comm). The basidiospores of A. sororcula are (8.8)9.7–12.8(16.8) x (8)8.8–12(15.5) µm (Tulloss et al 1992).
Amanita orientifulva Zhu L. Yang, M. Weiss et Oberw., sp. nov. FIGS. 13–16, 18
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Etymology: orientifulva is proposed because of the eastern Asian mushroom’s similarity to A. fulva.
Basidiocarps (FIG. 13) medium-size to large. Pileus 5–14 cm diam, at first hemispherical, then convex to planoconvex, obtusely umbonate, fulvous to brown or dark brown (Raw Umber, Antique Brown, Cinnamon-Brown, Orange-Cinnamon, 5C6, 5D6–7) over disk, becoming ochreous, yellowish brown to fulvous (Ochraceous-Tawny to Ochraceous-Orange, 5B5–6, 5C6, 4A4–5) toward margin, often with an indistinctly darker-colored ring-like zone at the proximal end of the marginal striation, glabrous; margin tuberculatestriate (0.2–0.35R, rarely up to 0.5R), nonappendiculate; trama white to dirty white to cream-colored, unchanging. Lamellae free, crowded, white to cream-colored (Cream Color, 1A1–2), with brown to brownish edges; lamellulae truncate, plentiful, evenly distributed. Stipe 8–15(–20) x 0.5–3 cm, usually tapering upward, with apex slightly expanded, dirty white to brownish (1A1, much paler than 5B4), covered with brown to fulvous or grayish brown (5B3–5) squamules; context white, hollow in center; basal bulb lacking; volva saccate, (2–)4–6(–9) x (1–)1.5–5 cm, membranous, outer surface white (1A1), often with rusty (6C5–6) spots, upper margin often brownish, inner surface pale brown (paler than 5B4), 1–3 mm thick, with limbus internus placed rather high on inside of volval limb or at point of attachment between stipe and volval limb. Annulus lacking. Odor indistinct.
Lamellar trama bilateral. Mediostratum 40–50 µm wide, composed of fairly abundant subfusiform to ellipsoid or clavate, inflated cells (45–100 x 15–25 µm); filamentous hyphae abundant, 3–7 µm wide; vascular hyphae rare. Lateral stratum composed of abundant clavate to subfusiform, inflated cells (55–100 x 20–30 µm), diverging at an angle of 30–45° to mediostratum; filamentous hyphae abundant, 2–7(–12) µm wide. Subhymenium (FIG. 14) 30–50 µm thick, with 2–3(–4) layers of subglobose to ovoid or short ellipsoid cells, 10–25(–30) x 10–20(–25) µm. Basidia (FIG. 14) 50–85(–95) x 15–20(–22) µm, clavate, 4-spored; sterigmata 5–8 µm long; basal septa without clamps. Basidiospores (FIG. 15) (585/27/17) (8.5–)10.0–14.0(–17.0) x (8.0–)9.5–13.0(–16.5) µm (Q = 1.0–1.12[–1.23], Q = 1.06 ± 0.04), globose to subglobose, rare broadly ellipsoid, inamyloid, colorless, hyaline, thin-walled, smooth; apiculus small. Lamellar edge microscopically appearing as a sterile, somewhat gelatinized, yellowish strip 100–200 µm wide in side view, composed of very abundant, ovoid to subglobose or sphaeropedunculate, thin-walled, colorless, inflated cells (25–70 x 20–40 µm), sometimes showing yellowish contents and appearing single and terminal or in terminal chains of 2–3; filamentous hyphae abundant, 2–5(–8) µm wide, gelatinized, colorless, hyaline or yellowish, ± running parallel to the lamellar edge. Pileipellis 60–100 µm thick; upper layer (30–50 µm thick) strongly gelatinized, composed of ± radially arranged, 1–3 µm wide filamentous hyphae, thin-walled, colorless to subcolorless; lower layer (30–50 µm thick) composed of radially and compactly arranged, 3–8(–12) µm wide filamentous hyphae, often with brownish to brown, vacuolar pigments; vascular hyphae rare (to scattered), 3–12 µm wide. Interior of volval limb on stipe base composed of two layers intergrading into each other. Outer layer (FIG. 16) composed of longitudinally to irregularly arranged elements: filamentous hyphae dominant, (2–)4–10(–12) µm wide, colorless, hyaline or sometimes with brownish to brown contents, slightly thick-walled (ca 0.5 µm thick), branching, anastomosing; inflated cells scattered to fairly abundant, subglobose to ovoid (40–70 x 30–55 µm) to long ellipsoid to subfusiform (80–90 x 30–35 µm), colorless, hyaline or with brownish contents, slightly thick-walled (ca 0.5 µm thick), usually single and terminal, sometimes in chains of 2; vascular hyphae scattered to locally conspicuous, 3–10 µm wide. Inner layer (FIG. 18) composed of very abundant 2–8 µm wide, colorless, hyaline, slightly thick-walled filamentous hyphae (up to 0.5 µm thick); inflated cells fairly abundant to locally abundant, subglobose to ovoid (35–100 x 30–85 µm), long ellipsoid to subfusiform (55–120 x 15–40 µm), single and terminal or intercalary, or in chains of 2–3, colorless, hyaline to subhyaline, thin- to slightly thick-walled (ca 0.5 µm thick); vascular hyphae rare, 3–10 µm wide. Outer surface of the volval limb similar to the interior of the outer layer but often with more abundant, somewhat gelatinized filamentous hyphae; inner surface strongly gelatinized, composed of 1.5–3 µm wide filamentous hyphae. Volval remnants on pileus lacking. Stipe trama composed primarily of longitudinally arranged, long clavate, terminal cells, 200–350 x 30–40 µm; filamentous hyphae 2–8 µm wide, scattered (in interior) to abundant (on stipe surface), colorless, hyaline (in interior) or with yellowish, vacuolar pigments (on stipe surface); vascular hyphae rare, 2–15 µm wide.
Habit, habitat, and distribution. – Solitary or scattered on soil under Abies, Quercus and/or Salix, sometimes under Castanopsis; fruiting from June to September in southwestern China at 1300–4200 m elevation.
Specimens examined..
CHINA. GUIZHOU PROVINCE: Suiyang County, Kuankuoshui, altitude 1300 m, 17 Jun 2000, X. L. Wu 819 (HKAS 35989). SICHUAN PROVINCE: Daocheng County, altitude 3800 m, 4 Jul 1998, Zhu L. Yang 1984 (HKAS 32531). Jiulong County, Jichoushan, altitude 3500 m, 11 Sep 1996, M. S. Yuan 2682 (HKAS 31133). Kangding County, Gonggashan, 14 Jul 1984, J. J. Su & H. A. Wen 1129 (HMAS 50987, as Amanita fulva in Ying et al 1994). Luding County, Moxi, altitude 3000 m, 12 Aug 1997, P. Q. Sun 2906 (HKAS 31360). Xiangcheng County, Daxueshan, altitude 4060 m, 24 Jul 1998, Zhu L. Yang 2423 (HKAS 32530); same locality, altitude 4000 m, 27 Jul 1998, Zhu L. Yang 2474 (HKAS 32470). Xiangcheng County, Reda, altitude 3450 m, 15 Jul 1998, Zhu L. Yang 2333 (HKAS 32483); same locality, altitude 3600 m, 16 Jul 1998, Zhu L. Yang 2344 (HKAS 32457). Xiangcheng County, Wumingshan, altitude 4100 m, 12 Jul 1998, Zhu L. Yang 2290 (HKAS 32478). YUNNAN PROVINCE: Deqin County, Meilishi, altitude 4200 m, 29 Aug 2000, Zhu L. Yang 3017 (HKAS 36604). Lijiang County, Laojunshan, altitude 3850 m, 13 Aug 2000, Zhu L. Yang 2911 (HKAS 36586). Yiliang County, Mugan, altitude 1800 m, 20 Sep 2000, Zhu L. Yang 2519 (HKAS 32664). Zhongdiang County, Daxueshan, altitude 3800 m, 24 Aug 2000, Zhu L. Yang 3016 (HKAS 36603). Zhongdiang County, Dongwang, altitude 3800 m, 26 Jul 1998, Zhu L. Yang 2461 (HKAS 32522, HOLO-TY PE); same locality, altitude 3800 m, 26 Jul 1998, Zhu L. Yang 2459 (HKAS 32464).
Commentary. –
Amanita orientifulva, belonging in Amanita subgenus Amanita section Vaginatae (Fr.) Quél. in the sense of Yang (1997), is rather common and widely distributed in China. This species is very similar to A. fulva (Schaeff.) Fr., originally described from Europe. In fact, A. orientifulva usually was regarded as A. fulva in China (Ying et al 1994). However, there are a few features separating A. orientifulva from A. fulva. The main morphological or anatomical difference between A. orientifulva and A. fulva might be the volval structure. The interior of the volval limb of A. fulva (GERMANY. BAVARIA: Bayerisches Wald, 18 Oct 1987, N. Arnold box II [L]) can be distinguished into two layers: The outer layer is composed of loosely and irregularly arranged, fairly abundant to abundant filamentous hyphae and fairly abundant to abundant inflated cells; the inner layer is composed predominantly of irregularly arranged filamentous hyphae, mixed with scattered to locally fairly abundant inflated cells (FIGS. 17, 19). Hyphae on the outer surface of the volva in A. fulva usually are broken during fruit body development. On the contrary, the outer layer of the volva of A. orientifulva is primarily composed of much more compactly and more or less longitudinally arranged, filamentous hyphae mixed with fewer inflated cells and scattered to locally conspicuous vascular hyphae, and the inner layer consists of more inflated cells (FIGS. 16, 18). Hyphae on the outer surface of the volva in A. orientifulva do not break during fruit body development. Furthermore, A. orientifulva is distinguished from A. fulva by its generally larger basidiocarps often with a somewhat darker-colored ring-like zone at the proximal end of the marginal striations of the pileus and somewhat larger basidiospores (Tulloss 2000). The basidiospores of A. fulva are (9.0–)10.0–12.5(–19.3) x (8.2–)9.3–12.0(–15.5) µm (Tulloss 2000).
Amanita orientifulva is phenetically somewhat similar to A. sampajensis Sathe et Kulkarni, A. fuligineodisca Tulloss, Ovrebo et Halling and A. humboldtii Singer. Amanita sampajensis, known from India, has an isabelline to hazel pileus, dark olivaceous brown stipe, white volva and a pileipellis with wavy hyphae with olivaceous contents (Sathe et al 1980, Tulloss et al 1992). Amanita fuligineodisca and A. humboldtii, both originally described from Colombia, have smaller fruit bodies, usually with longer marginal striations on the darker pileus and thinner pileipellis. Furthermore, there is no pronounced division of structure within the interior of the volval limb of A. fuligineodisca and A. humbolditii (Tulloss et al 1992, Tulloss pers comm).
In the protologue of A. aporema Boedijn, a poorly known species described from Indonesia, it was supposed that A. aporema might be confused with A. fulva (Boedijn 1951:320). We have studied the only specimen cited by Boedijn under A. aporema (INDONESIA. SUMATRA: Batang Paleopoeh, exact day unknown, Jul 1924, E. Jacobson s.n. [HOLOTY PE, BO]). It reveals that the basidiospores are inamyloid, (35/1/1) (9.0 –)9.5–11.0(–12.5) x (8.0 –)8.5–10.5 (–12.0) µm (Q = [1.0–]1.02–1.11[–1.15], Q = 1.08 ± 0.04) and clamps are common in all parts of the fruit body. These data indicate that A. aporema is more closely related to A. princeps Corner et Bas than to A. fulva and its allies. Based on the original description and examination of the poorly preserved holo-type, it appears that A. aporema differs from A. orientifulva at least by its dirty white, smooth stipe, somewhat smaller basidiospores and the presence of clamps.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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1 Corresponding author: Zhu Liang Yang, Kunming Institute of Botany, Chinese Academy of Sciences, Heilongtan, Kunming 650204, P.R. China. Fax: 0086-871-5150227; E-mail: zlyang{at}public.km.yn.cn; fungi{at}mail.kib.ac.cn
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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